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Spec Dinosauria: Ungulapedia
HISTORY The ancestry of the ungulapedes is traceable back to Anserodromeus, a small crested basal-hadrosauroid from Paleocene-Early Eocene Africa. Africa, alone amongst the Gondwanan landmasses, seems to have kept its duckbills after the K-Pg boundary. Despite a setback in the Late Eocene, the descendents of Anserodromeus flourished and diversified. While many forms became bulky giants, echoing the great herds of the Late Cretaceous, one lineage became agile woodland runners, gaining their distinctive hand-hoof during the Oligocene. With the gradual drying of the African continent as the Neogene progressed, the larger, more conservative hadrosaurs went into decline---to the boon of the early ungulapedes, which soon spread into the vacated niches. The Pliocene was a time of spectacular evolutionary innovation for the ungulapedes with the Afrohadrosauridae becoming the most numerous and diverse herbivores on the continent. When the long isolation of the African continent ended in the Miocene, great herds of ungulapeds marched across the Sinai into Eurasia. With efficient digestive systems, live-birth, long endurance, and a tightly knit herd structure, the invaders quickly made themselves at home, flourishing and diversifying at the expense of local herbivorous clades like the eurolophs. During the Pliocene, several lineages of African ungulapeds appear in European and south Asian deposits. These hadrosaurs seem to have driven the large eurolophids to extinction, and may also have contributed to the decline of small caenoceratopsians. However, when the Ice Ages began, only some coronolophids and catoblepids remained in southern Europe, while other forms were pushed north. During the interglacial periods, some new African saurolopines migrated to Europe, along with aquatic titanosaurs, but were again driven back or to extinction as the glaciers returned. Controversial evidence suggest that a few made it to North America, but these wanderers did not survive for long. The ice ages eliminated many species but thanks to the continued speciation of the adaptable ealine and catoplebine formosicorns, the ungulapeds have been able to maintain their numbers in Eurasia, and are today one of the "big three" successful large herbivore clades in the region along with the ceratopsians and sauropods. BIOLOGY Ungulapedes are most easily recognized by their forefeet. Digits 2 and 3 have become fused and the combined unguals protrude from the flesh to form a single large hoof. Digit 4 is reduced and completely internalized however, the 5th digit remains free in all forms except the elumbe and often bears a large claw. The most remarkable aspect of these hadrosaurs is not to be found in their feet, however, but involves their reproductive habits. With the exception of the archaic paleungulapodes, the majority of the Ungulapedia has largely or totally abandoned nesting altogether and carries their incubating offspring with them. Most female ungulapedes possess a large, highly elastic gular pouch that can be partially sealed off from the rest of the mouth via muscular contractions. These broodpouches are used for egg-storage and are both highly padded with rubbery tissue both for insulation and to prevent damage to the egg. The mother ungulapede lays one or two eggs that vary from between 15 to 45 cm in length. Once deposited on the ground, the egg is taken into her mouth and gently manoeuvered into the broodpouch. Most species lay either single eggs or produce two but discard or consume one of them. These forms have a simple, single-chambered broodpouch. Some ealines, however will lay and nurture two eggs, their broodpouch having evolved a bifurcated, two-pocketed configuration (bearing a passing resemblance to a mammalian scrotum) allowing them to safely transport both eggs. With her broodpouch fully laden, the mother is still capable of eating and drinking albeit in a somewhat slow and awkward manner. Sometimes she will remove the egg from her broodpouch and place it on the ground or a brief period while she gorges. Incubation usually takes between 1 and 4 months depending on the species and local conditions. Upon sensing that hatching is imminent, the mother coughs up the egg and deposits on the ground, sometimes assisting the youngster by breaking the shell herself. Once it finds its feet, the large hatchling is fully capable of fending for itself although the majority of ungulapede mothers (and some ealine fathers) provides additional protection to their offspring after hatching. The young of highly gregarious species leave their mother's side in a few days or weeks to form juvenile cr?hes, which either head off on their own or stay to enjoy the protection of the entire herd. Those of less gregarious forms tend to stay with their parents for longer periods. Regardless of parental care , the young feed ravenously and can put on as much as 40 kg in their first month. Most reach sexual maturity in between 2-4 years. This unique breeding strategy confers many advantages. For most dinosaurs, the nesting period is a dangerous time for both adults and offspring as their mobility is severely restricted. There is the ever-present danger of predators, egg-thieves and natural disasters befalling the nest site. By dispensing with the nest, the ungulapedes have left behind many of these problems at the expense of reduced clutch size. A female with a full broodpouch can still feed, drink, evade predators, tend to her previous generation of offspring or even mate and thus get the next egg ready once the one in the throat hatches. These benefits become even more vital on the great plains of Africa and Asia where the ungulapedes thrive. Here, vast herds must wander great distances as they follow highly seasonal and irregular food supplies. With this sort of lifestyle, having to spend several months tied down to a static nesting site would be a severe handicap. The development of this brood-pouch is probably responsible for the ungulapeds' tremendous sucsess in Africa and Eurasia, where they are they are by far the dominant clade of small herbivores. Indeed, only their intolerance to cold, it seems, has prevented the ungulapeds from spanning the globe. PALEOUNGULAPODOIDAE (Blowhards and oopas) Generally small and completely restricted to southern and central Africa, the paleoungulapods (confusingly it's "pods" not "pedes") are the most primitive extant members of Ungulapedia. The cape blowhard is a primtive ungulapede that lives in open dry plains of Southern Africa. It feeds on many plant species, eating grass, fruit, tubers and leaves. It is generally solitary, declaring its territory with its distinctive honking call and a comical-looking nasal balloon. Cape blowhard, Erythronasua srnkai ''(southern Africa) Spec's Congo harbor a number of oddities, relic species from a time when Africa was a lush continent, covered with jungles of which the present greenery is merely a fragment. One of these relics is the primitive oopa, a paleoungulapod most closely related to the blowhards of Africa's southern coast. Like their smaller cousins, oopas lack horns, sporting enlarged brow ridges and a laterally flattened nasal crest in their stead. These timid herbivores are solitary in their habits, although young calves with trail behind their mother for up to a year before heading off on their own. Males, larger than females and more brightly-colored, can be quite ill-tempered and deliver vicious kicks with their front hooves to any creature stupid or unlucky enough provoke them. Oopa, ''Cranitus cristatus ''(central Africa) AFROHADROSAURIDAE (Saurolopes, lanceheads, and bowhorns) The saurolope body-type. Note the hooved front feet, long snout, and six squamosal horns. About one half of the ungulapedes belong to the family Afrohadrosauridae (the saurolopes). They are largely restricted to Africa with a few species in Asia Minor and India. South of the Sahara, vast nomadic herds of brightly coloured saurolopes blanket the landscape. Afrohadrosaurids are generally fairly gracile animals with long square-tipped snouts. Most species have between 2-6 narrow squamosal horns but lack any sort of cranial or nasal ornamentation. All saurolopes are at least partial grazers, none survive on a completely leafy diet. The six-horned saurolope is a creature of scrub and savannah; rarely far from water although it is not closely associated with it. It lives in herds of up to 30 individuals led by an elderly bull with young males forming separate herds of the same size. They feed on grasses and other vegetation. Six-horned saurolope, ''Saurolopus sexicornis ''(sub-Saharan Africa) 5 species of ''Oryxosaurus can be found throughout Africa. The green lancehead is found across the eastern savannas, corresponding to our world's Eritrea-to-northern-Angola. It lives in herds of 10-50 individuals, often associating with sauropods or other saurolope species, and feeds on grass and leaves. Green lanceheads can go for long periods without water. Green lancehead, Oryxosaurus chloris ''(Eastern sub-Saharan Africa) The only member of its genus, the bowhorn is a rather odd saurolope, allied with the lanceheads (genus ''Oryxosaurus), but distinguished by its branching squamosal horns (as in the megacornid hornmeister). Along with this ornamentation, bowheads also sport a pair of smaller, triangular horns in the region of the cheeks. Unlike most other saurolopes, bowhorns are solitary creatures, mostly preferring to browse near rivers and forest margins by themselves or in small groups. During the mating season, however, just before the rains fall, large groups of bowhorns may gather together, males displaying their fantastic branching horns and brilliant orange and blue facial markings. Bowhorn, Quadricornis magnificus ''(Eastern sub-Saharan Africa) Lowlands angarang Tonitrobalus fracanaulon notios (sub-Saharan Africa) Comparison of the plains angarang and the closely related whiffleboom The doracasaurs are the fastest and smallest of the saurolopes. They seldom exceed 2.5 meters in length, and are known to attain speeds acomparable to those of springbok and American pronghorn antelopes. Doracasaurs are mixed browser-grazers, and are notoriously skittish, and outrun all but the fastest abelisaurian predators. Doracasaur, ''Doracasaurus racilis ''(Southern Africa) The bristlehorn, largest of all saurolopes, rivals the great hormeister and elumbe in size, and is quite dangerous. These large -bodied marshdwellers are generally solitary, shunning their own kind out of breeding season. They are sometimes seen browsing alongside herds of mokele for protection. Bristlehorn, ''Xenocornis major ''(sub-Saharan Africa) CIRAPHADRIDAE (Cirafs) Not as widespread or specious as the related afrohadrosaurids, the ciraphadrids are somewhat more diverse. Occupying both moose-like, swamp-dwelling forms, as well as deep-forest browsers and the familiar high browser of the savannah, the evolutionary success of the ciraphadrids is indisputable. This creature, recenly discovered in Africa's Congo Basin, and named only "ciraphid" bears a close resemblance to the probable generalized browsing ancestor of Ciraphidae. The ciraf, African high-browser of the smallest type, is often dwarfed by the titanosaur giants with which it lives. In the Africa of our home timeline, however, this 7.5 m long ungulaped would be one of the largest animals of the savanna. Ciraf, ''Ciraphadrus longicervix ''(south-Saharan Africa) ULTRACORNIDAE (Hornmeisters) This monotypic family contains the giant browsing hornmeister, the only ungulapede to combine a unicorn-spike and four squamosal horns. Fossil ultracornids both larger and smaller than the extant species are known from Mio-Pleistocene Africa and Eurasia. Phylogenetically, the hornmeister falls midway between the Afrohadrosauridae and the formosicorns, and seems to represent an early sidebranch on the evolutionary line to Formosicorna. The largest African ornithischian, this 10-meter animal is still dwarfed by the giant grassbags with which it often associates. A creature of savanna and and open forests throughout eastern and southern Africa, the hornmeister lives in large herds, usually of around 30-50 individuals but occasionally up to 200. Old males tend to be solitary. It feeds on leaves and tender branches and sometimes digging for tubers. Hornmeister, ''Ultracornis brenseni ''(south-Saharan Africa) Hornmeister FORMOSICORNIDAE (Elumbes, orths, catoblepines, and ealines) Formosicornidae is a largely Eurasian a ungulapede radiation somewhat related to Ultracornidae. The three branches of Eurasian Formosicornidae to scale. From left to right: yale (ealine), majestic orth (coronolophid), and bruton (catoblepine). The taxonomy of the formosicorns has been a nightmare of duplicity and classificatory juggling with different species grouped in the families "Boreosauridae" (no longer used), Cornucantidae, Ultracornidae (in part), Afrohadrosauridae (in part), Catoblepidae, Ealidae and Tricornidae. At one point, the formicornids were even given their own sub-class, Formosicornia, ridiculous as that may sound. Recent DNA-hybridization data and exhaustive anatomical and palaeontological studies have shown strong support for the monophyly of these seemingly disparate forms in a single family-level clade, Formosicornidae. Formosicorns differ from the African ungulapeds in having larger nasal cavities, the presence of circumnarial depressions and a maximum of 2 squamosal horns. They and the African hornmeister share a distinctive "unicorn horn", a spikey protruberance on the forehead whose horny base is formed by a fusion of the frontals (and sometimes the nasals) into a rugose lump. Formosicorns generally exhibit a lesser degree of sexual dimorphism than the other large ungulapeds, with both sexes often sporting spectacular horns. With the exception of some catoblepids, the formosicorns do not form herds of the same as magnitude as the African saurolopes. Formosicorns originated in Africa from basal "proafrohadrosaur" stock in the Early Miocene. While their afrohadrosaurid cousins became grazers, the early formosicorns stayed fairly generalised and were amongst the first species "off the boat" when the link with Asia was formed. They proved to be the most adaptable of the invading ungulapeds and soon diverged into dozens of species. With the exception of the bulky grazing cornucantids, most of these were lightly built browsers living in the shadow or the now extinct giant afrohadrosaurs and ultracornids. With the onset of the Ice Ages, the formosicorns proved to be better able to handle the cooler conditions than the their cousins. In a very short space of time, the family underwent another explosive diversification event producing a host of new Pleistocene forms including the armoured grazing catoblepines. While they lost ground to the saurolopes in their ancestral African homeland, the formosicorns wasted no time in becoming the dominant Eurasian ungulaped clade. The last glaciation period hit Europe's ornithischians hard, driving the last of the local ultracornids, saurolopes and cornucantines to extinction. The modern ungulaped assemblage largely repopulated Europe from other parts of Eurasia. Such immigrants include the tricorns and catoblepines (which extended their annual migrations to Southern Europe even during the glaciation, but didn't spend the winters there.) Since the climate has again started to get colder predicting a new cycle of glaciation, the hadrosaurs are no longer found north of the Baltic. 6000 year old bone findings however prove that catoblepines once roamed near the arctic circle. CORNUCANTINAE (Elumbes) In Africa, with the afrohadrosaurid saurolopes now dominating most of the large herbivore guilds they are restricted to the giant elumbe, a few small ealids on the North coast and a some plesiomorphic old-endemic forest forms. Elumbe The elumbe is a recent immigrant from Europe, the largest and last survivor of the cornucantidae, basal formosicorn behemoths (closely allied to the catoblepids) that flourished in Plio-Pleistocene western Eurasia, but succumbed to extinction in Europe during the last Ice Age. At up to 8.5 m in length and weighing in at a little under 7 tonnes, it is by far the most massive of the ungulapeds. Majestic orth, Cornucantus belli'' (Central and southern Africa) Notable features of the elumbe include the loss of digit 5, the enormous "unicorn-horn" whose base is formed from the fusion of the frontals and posterior nasals, equal-sized pedal digits (the middle toe is enlarged in extant Eurasian formosicorns) with relatively small unguals and reduced squamosal horns. It feeds primarily on grasses but will also take some leaves and tubers. It also drinks a lot, staying close to reliable water sources. The elumbe is a solitary, territorial animal, coming together only to mate. The single calf is raised by the mother alone. They are nearsighted and bad-tempered, charging at anything that moves. It was an elumbe that was responsible for crippling the large male molok which was later killed by the thebirds - suffice to say that even had the molok not encountered the cityfinch colony, it would not have been making any more molokettes. CORONOLOPHIDAE (Orths) The diversity of horn shapes among orths, beginning from top left: cuinocco, dwarf orth, dhar, majestic orth (female), brass orth. (The shape of the female's horn is drawn beside the head of the male) The coronolophids, or orths, were the first group of formosicorns the split off as the clade migrated into Europe. These creatures are relatively primative, with rather weak jaw muscles that restrict their diets to tender leaves and shoots. Despite their gastronomic limitations, orths are quite common in western-central Eurasia, with populations extending north to Fennoscandia and the British Isles. These herbivores are larger than most ealines (though not so large as the catoblepines) and are distinguished by their large, flattened central horn (morpholgically the same as the ealines' "unicorn horn") which is covered with skin. Most orths hibernate during the winter. The 8 m long majestic orth is a prime example of the typically European hadrosaurs, the orths. The males' large crest-like central horn is covered with skin which is brightly coloured during the mating season. Males are also larger and more robust than females. Orth males gather a harem of females which they guard from veldraks and other males. Majestic orth, Coronolophus rex (Central Europe and Asia) Majestic orth (male) in rut Almost as large as a majestic orth at 7.5 meters, but probably more closely related to the cuinocco, the photorth is is a beautiful and secretive denizen of the jungles of south and southeast Asia. For predators, the photorth is not an easy prey. Male photorths have three horns on their head, while female sport one large and sharp unicorn horn in the place of an orth's normal decretive crest. The males are aggressive during rut, but the females, with their sharp unicorn horns, are lethal. Photorth, Photor sappimus (South and Southeast Asia) The cuinocco is the largest and most secretive of the formosicorns, dwells on the Scandinavian peninsula and the British Isles. Denizens of the deep forest, cuinoccos are only seen singly, or in mated pairs. Like many northern formosicorns, cuinoccos hibernate through the winter in caves or in riverside burrows dug with spade-like front hooves. These creatures may still be seen in winter, however, as they must rouse themselves, periodically, to forage for food. Cuinocco,Cervicerosaurs mirabilis (Scanindavia and British Isles) CATOBLEPINAE (Gonnucs and beluboses) The catoblepines are the largest Eurasian ungulapedes. Like their cousins, the ealines, they are primarily grazers which often gather in huge migratory herds. In contrast to the graceful running ealines, however, the catoblepines have become so massive that they cannot manage anything more strenuous than a brisk amble. The middle toe of catoblepines is not as enlarged as those of ealines with no significant reduction of the other toes. The unicorn horn is generally either a short spike or a flattened boss. The horn-base in catoblepines is formed by the fusion of both the frontals and the posteriormost nasal bones. Though their large size and horns already make them formidable adveseries, catoblepines also possess armour protection in the form of numerous small osteoderms. The first catoblepines seem to appear in Asia Minor during early Pliocene, and probably evolved from primitive ealines. Later they evolved into humongous sizes, taking over the niche of the eurolophine rhinolophs. Since the climate has again started to get colder predicting a new glaciation period, the hadrosaurs are no longer found north of the Baltic. 6000 year old bone findings however prove that catoblepines once roamed near the arctic circle. The Indian catoblepine, the gonnuc, hasn't grown nearly as big as its European relatives, because ceratopsians still dominate the large herbivore niches in Southern asia. It lives in small, loose herds, where the dominance among males is determined by butting contests. As a result, the gonnuc's central horn has evolved into a large, flat boss. The beluboses aren't as migratory as most of their close relatives, though they also roam around the grasslands of south and central Europe. Gonnuc, Bonnacon malleocranium (South Asia) and Belubos, Catoblepas bucinator (Central Eurasia) Head of gonnuc Large herds of brutons roam across central and eastern Europe, spending the summers near the southern shores of the Baltic sea or the edge of the Russian taiga and returning south for the winter. Bruton, Catoblepas migratoris (Western Eurasia) Grazing bruton EALINAE (Yales and kirin) Often referred to as tricorns, ealines are the most widespread and speciose ornithopods in Eurasia (about 50 species). They are distinctive in possessing an enlarged middle toe on the pes with a reduction of the other two toes, a long unicorn-horn whose base is restricted to the frontals. Most are grazers that live in pairs or small to moderate sized herds. Unlike the polygamous catoblepines, most ealide species form stable pair or trio bonds. Not surprisingly, the tricorns exhibit conspicuous external sexual dimorphism. The Eurasian yales are quite similiar to their distant cousins, the african saurolopes. These lightweight hadrosaurs are fast gallopers, which live in herds that follow the migratory brutons. There are also several yale species living in the middle east and India. Yale, Eale recticornis (Western Eurasia) Kirins are the yales of eastern Asia. These antelope-like ealinids live in herds of about 10-30 individuals and ruled by an old male. The long, straight central horn of the kirin is a formidable weapon, but mostly used only for intimidation between males or against predators. Only kirin males have a long, forward-pointing horn. The female's horn is shorter and points straight up from the skull. The horn structure is remarkably similar to giant African elumbe, though it seems to be mere convergence. Kirin, Kirin chilin (Eastern Eurasia) ''Brian Choo, Matti Aumala, Daniel Bensen ''and ''Ville Sinkkonen '' ,=Erythronasua srnkai (Cape Blowhard) ,=Paleoungapodoidae=| | `=Cranitus cristatus (Oopa) =Ungalpedia=| | ,=Saurolopus sexicornis (Six-horned Saurolope) | ,Afrohadrosauridae=| | | | ,=Oryxosaurus chloris (Green Lancehead) | | `=| | | `=Quadrocornis Magnificus (Bowhorn) | ,=| | | `=Cirphadrididae=Ciraphadrus longicervix (Ciraf) `=| | ,=Ultracornidae=Ultracornis benseni (Hornmeister) `=| / / ,-------' / ,=Cornucantinae=Cornucantus belli (Elumbe) / | | | \=Formosicornidae=| / ,=Coronolophus rex (Majestic orth) / ,=| / | | ,=Photor sappimus (Photorth) / | `=| / | `=Cervicosaurus mirabilis (Cuinocco) `=Coronolophidae=| | ,=Bonnacon malleocranium (Gonnuc) | ,=Catoblepinae=| | | | ,=C. bucinator (Belubos) `=| `=Catoblepas=| | `=C. migratoris (Bruton) | | ,=Eale recticornis (Yale) `=Ealinae=| `=Kirin chilin (Kirin) Category:Spec Dinosauria Category:Reptiles Category:Animals Category:Dinosaurs